Prediction and confirmation of seasonal migration of Pacific sardine (Sardinops sagax) in the California Current Ecosystem

During the last century, the population of Pacific sardine (Sardinops sagax) in the California Current Ecosystem has exhibited large fluctuations in abundance and migration behavior. From approximately 1900 to 1940, the abundance of sardine reached 3.6 million metric tons and the “northern stock” migrated from offshore of California in the spring to the coastal areas near Oregon, Washington, and Vancouver Island in the summer. In the 1940s, the sardine stock collapsed and the few remaining sardine schools concentrated in the coastal region off southern California, year-round, for the next 50 years. The stock gradually recovered in the late 1980s and resumed its seasonal migration between regions off southern California and Canada. Recently, a model was developed which predicts the potential habitat for the northern stock of Pacific sardine and its seasonal dynamics. The habitat predictions were successfully validated using data from sardine surveys using the daily egg production method; scientific trawl surveys off the Columbia River mouth; and commercial sardine landings off Oregon, Washington, and Vancouver Island. Here, the predictions of the potential habitat and seasonal migration of the northern stock of sardine are validated using data from “acoustic–trawl” surveys of the entire west coast of the United States during the spring and summer of 2008. The estimates of sardine biomass and lengths from the two surveys are not significantly different between spring and summer, indicating that they are representative of the entire stock. The results also confirm that the model of potential sardine habitat can be used to optimally apply survey effort and thus minimize random and systematic sampling error in the biomass estimates. Furthermore, the acoustic–trawl survey data are useful to estimate concurrently the distributions and abundances of other pelagic fishes.

Life-stage recruitment models for Atlantic cod (Gadus morhua) and haddock (Melanogrammus aeglefinus) on Georges Bank

Ichthyoplankton surveys have been used to provide an independent estimate of adult spawning biomass of commercially exploited species and to further our understanding of the recruitment processes in the early life stages. However, predicting recruitment has been difficult because of the complex interaction of physical and biological processes operating at different spatial and temporal scales that can occur at the different life stages. A model of first-year life-stage recruitment was applied to Georges Bank Atlantic cod (Gadus morhua) and haddock (Melanogrammus aeglefinus) stocks over the years 1977–2004 by using environmental and densitydependent relationships. The best lifestage mortality relationships for eggs, larvae, pelagic juveniles, and demersal juveniles were first determined by hindcasting recruitment estimates based on egg and larval abundance and mortality rates derived from two intensive sampling periods, 1977–87 and 1995–99. A wind-driven egg mortality relationship was used to estimate losses due to transport off the bank, and a wind-stress larval mortality relationship was derived from feeding and survival studies. A simple metric for the density-dependent effects of Atlantic cod was used for both Atlantic cod and haddock. These life stage proxies were then applied to the virtual population analysis (VPA) derived annual egg abundances to predict age-1 recruitment. Best models were determined from the correlation of predicted and VPA-derived age-1 abundance. The larval stage was the most quantifiable of any stage from surveys, whereas abundance estimates of the demersal juvenile stage were not available because of undersampling. Attempts to forecast recruitment from spawning stock biomass or egg abundance, however, will always be poor because of variable egg survival.

Stage-specific vertical distribution of Alaska plaice (Pleuronectes quadrituberculatus) eggs in the eastern Bering Sea

The stage-specific distribution of Alaska plaice (Pleuronectes quadrituberculatus) eggs in the southeastern Bering Sea was examined with collections made in mid-May in 2002, 2003, 2005, and 2006. Eggs in the early stages of development were found primarily offshore of the 40-m isobath. Eggs in the middle and late stages of development were found inshore and offshore of the 40-m isobath. There was some evidence that early-stage eggs occur deeper in the water column than late-stage eggs, although year-to-year variability in that trend was observed. Most eggs were in the later stages of development; therefore the majority of spawning is estimated to have occurred a few weeks before collection—probably April—and may be highly synchronized among local spawning areas. Results indicate that sampling with continuous underway fish egg collectors(CUFES) should be supplemented with sampling of the entire water column to ensure adequate samples of all egg stages of Alaska plaice. Data presented offer new information on the stage-dependent horizontal and vertical distribution of Alaska plaice eggs in the Bering Sea and provide further evidence that the early life history stages of this species are vulnerable to near-surface variations in hydrographical conditions and climate forcing.

Larval abundance of summer flounder (Paralichthys dentatus) as a measure of recruitment and stock status

Summer flounder (Paralichthys dentatus) is one of the most economically and ecologically important estuarine-dependent species in the northeastern United States. The status of the population is currently a topic of controversy. Our goal was to assess the potential of using larval abundance at ingress as another fishery independent measure of spawning stock biomass or recruitment. Weekly long-term ichthyoplankton time series were analyzed from Little Egg Inlet, New Jersey (1989–2006) and Beaufort Inlet, North Carolina (1986–2004). Mean size-at-ingress and stage were similar between sites, whereas timing of ingress and abundance at ingress were not similar. Ingress primarily occurred during the fall at Little Egg Inlet and the winter at Beaufort Inlet. These findings agree with those from earlier studies in which at least two stocks (one north and one south of Cape Hatteras) were identified with different spawning periods. Larval abundance at Little Egg Inlet has increased since the late 1990s and most individuals now enter the estuary earlier during the season of ingress. Abundance at Little Egg Inlet was correlated with an increase in spawning stock biomass, presumably because spawning by larger, more abundant fish during the late 1990s and early 2000s provided increased larval supply, at least in some years. Larval abundance at ingress at Beaufort Inlet was not correlated with spawning stock biomass or with larval abundance at ingress at Little Egg Inlet, further supporting the hypothesis of at least two stocks. Larval abundance at Little Egg Inlet could be used as a fishery-independent index of spawning stock size north of Cape Hatteras in future stock assessments. Larval occurrence at Beaufort Inlet may provide information on the abundance of the stock south of Cape Hatteras, but additional stock assessment work is required.

Biomass and reproduction of Pacific sardine (Sardinops sagax) off the Pacific northwestern United States, 2003–2005

The Pacific sardine (Sardinops sagax) is distributed along the west coast of North America from Baja California to British Columbia. This article presents estimates of biomass, spawning biomass, and related biological parameters based on four trawl-ichthyoplankton surveys conducted during July 2003 –March 2005 off Oregon and Washington. The trawl-based biomass estimates, serving as relative abundance, were 198,600 t (coefficient of variation [CV] = 0.51) in July 2003, 20,100 t (0.8) in March 2004, 77,900 t (0.34) in July 2004, and 30,100 t (0.72) in March 2005 over an area close to 200,000 km2. The biomass estimates, high in July and low in March, are a strong indication of migration in and out of this area. Sardine spawn in July off the Pacific Northwest (PNW) coast and none of the sampled fish had spawned in March. The estimated spawning biomass for July 2003 and July 2004 was 39,184 t (0.57) and 84,120 t (0.93), respectively. The average active female sardine in the PNW spawned every 20–40 days compared to every 6–8 days off California. The spawning habitat was located in the southeastern area off the PNW coast, a shift from the northwest area off the PNW coast in the 1990s. Egg production in off the PNW for 2003–04 was lower than that off California and that in the 1990s. Because the biomass of Pacific sardine off the PNW appears to be supported heavily by migratory fish from California, the sustainability of the local PNW population relies on the stability of the population off California, and on local oceanographic conditions for local residence.

Seasonal variability in ichthyoplankton abundance and assemblage composition in the northern Gulf of Mexico off Alabama

Multiyear ichthyoplankton surveys used to monitor larval fish seasonality, abundance, and assemblage structure can provide early indicators of regional ecosystem changes. Numerous ichthyoplankton surveys have been conducted in the northern Gulf of Mexico, but few have had high levels of temporal resolution and sample replication. In this study, ichthyoplankton samples were collected monthly (October 2004–October 2006) at a single station off the coast of Alabama as part of a long-term biological survey. Four seasonal periods were identified from observed and historic water temperatures, including a relatively long (June–October) “summer” period (water temperature >26°C). Fish egg abundance, total larval abundance, and larval taxonomic diversity were significantly related to water temperature (but not salinity), with peaks in the spring, spring–summer, and summer periods, respectively. Larvae collected during the survey represented 58 different families, of which engraulids, sciaenids, carangids, and clupeids were the most prominent. The most abundant taxa collected were unidentified engraulids (50%), sand seatrout (Cynoscion arenarius, 7.5%), Atlantic bumper (Chloroscombrus chrysurus, 5.4%), Atlantic croaker (Micropogonias undulatus, 4.4%), Gulf menhaden (Brevoortia patronus, 3.8%), and unidentified gobiids (3.6%). Larval concentrations for dominant taxa were highly variable between years, but the timing of seasonal occurrence for these taxa was relatively consistent. Documented increases in sea surface temperature on the Alabama shelf may have various implications for larval fish dynamics, as indicated by the presence of tropical larval forms (e.g., fistularids, labrids, scarids, and acanthurids) in our ichthyoplankton collections and in recent juvenile surveys of Alabama and northern Gulf of Mexico seagrass habitats.

Development and growth of hatchery-reared larval Florida pompano (Trachinotus carolinus)

Although the Florida pompano (Trachinotus carolinus) is a prime candidate for aquaculture, the problematic production of juveniles remains a major impediment to commercial culture of this species. In order to improve the understanding of larval development and to refine hatchery production techniques, this study was conducted to characterize development and growth of Florida pompano from hatching through metamorphosis by using digital photography and image analysis. Newly hatched larvae were transparent and had a large, elongate yolk sac and single oil globule. The lower and upper jaws as well as the digestive tract were not fully developed at hatching. Rotifers were observed in the stomach of larvae at three days after hatching (DAH), and Artemia spp. were observed in the stomach of larvae at 14 DAH. Growth rates calculated from total length measurements were 0.22 ±0.04, 0.23 ±0.12, and 0.35 ±0.09 mm/d for each of the larval rearing trials. The mouth gape of larvae was 0.266 ±0.075 mm at first feeding and increased with a growth rate of 0.13 ± 0.04 mm/d. Predicted values for optimal prey sizes ranged from 80 to 130 μm at 3 DAH, 160 to 267 μm at 5 DAH, and 454 to 757 μm at 10 DAH. Based on the findings of this study, a refined feeding regime was developed to provide stage- and size-specific guidelines for feeding Florida pompano larvae reared under hatchery con

Advances in methods for determining fecundity: application of the new methods to some marine fishes

Estimation of individual egg production (realized fecundity) is a key step either to understand the stock and recruit relationship or to carry out fisheries-independent assessment of spawning stock biomass using egg production methods. Many fish are highly fecund and their ovaries may weigh over a kilogram; therefore the work time can be consuming and require large quantities of toxic fixative. Recently it has been shown for Atlantic cod (Gadus morhua) that image analysis can automate fecundity determination using a power equation that links follicles per gram ovary to the mean vitellogenic follicular diameter (the autodiametric method). In this article we demonstrate the precision of the autodiametric method applied to a range of species with different spawning strategies during maturation and spawning. A new method using a solid displacement pipette to remove quantitative fecundity samples (25, 50, 100, and 200 milligram [mg]) is evaluated, as are the underlying assumptions to effectively fix and subsample the ovary. Finally, we demonstrate the interpretation of dispersed formaldehyde-fixed ovarian samples (whole mounts) to assess the presence of atretic and postovulatory follicles to replace labor intensive histology. These results can be used to estimate down regulation (production of atretic follicles) of fecundity during maturation.

All their eggs in one basket: a rocky reef nursery for the longnose skate (Raja rhina Jordan & Gilbert, 1880) in the southern California Bight

Skates (family Rajidae) are oviparous and lay tough, thick-walled eggs. At least some skate species lay their eggs in spatially restricted nursery grounds where embryos develop and hatch (Hitz, 1964; Hoff, 2007). After hatching, neonates may quickly leave the nursery grounds (Hoff, 2007). Egg densities in these small areas may be quite high. As an example, in the eastern Bering Sea, a site <2 km2 harbored eggs of Alaska skate (Bathyraja parmifera) exceeding 500,000/km2. All skate nursery grounds have been identified over soft sea floors (Lucifora and García, 2004; Hoff, 2007).

A nursery site of the Alaska skate (Bathyraja parmifera) in the eastern Bering Sea

A nursery site for the Alaska skate (Bathyraja parmifera) was sampled seasonally from June 2004 to July 2005. At the small nursery site (~2 km2), located in a highly productive area near the shelf-slope interface at the head of Bering Canyon in the eastern Bering Sea, reproductive males and females dominated the catch and neonate and juvenile skates were rare. Seasonal samples showed summertime (June and July) as the peak reproductive time in the nursery although some reproduction occurred throughout the year. Timeseries analysis of embryo length frequencies revealed that three cohorts were developing simultaneously and the period of embryonic development was estimated at 3.5 years and average embryo growth rate at 0.2 mm/day. Estimated egg case deposition occurred mainly during summertime and hatching occurred during winter months. Protracted hatching times may be common for oviparous elasmobranch species and may be directly correlated with ambient temperatures as evident from a meta-data analysis. Evidence indicates that the Alaska skate uses the eastern Bering Sea outer continental shelf region for reproduction and the middle and inner shelf regions as habitat for immature and subadults. Skate nurseries may be vulnerable to disturbances because they are located in highly productive areas and because embryos develop slowly.

Ichthyoplankton community in the Columbia River plume off Oregon: effects of fluctuating oceanographic conditions

Ichthyoplankton samples were collected at approximately 2-week intervals, primarily during spring and summer 1999−2004, from two stations located 20 and 30 km from shore near the Columbia River, Oregon. Northern anchovy (Engraulis mordax) was the most abundant species collected, and was the primary species associated with summer upwelling conditions, but it showed significant interannual and seasonal fluctuations in abundance and occurrence. Other abundant taxa included sanddabs (Citharichthys spp.), English sole (Parophrys vetulus), and blacksmelts (Bathylagidae). Two-way cluster analysis revealed strong species associations based primarily on season (before or after the spring transition date). Ichthyoplankton abundances were compared to biological and environmental data, and egg and larvae abundances were found to be most correlated with sea surface temperature. The Pacific Decadal Oscillation changed sign (from negative to positive) in late 2002 and indicated overall warmer conditions in the North Pacific Ocean. Climate change is expected to alter ocean upwelling, temperatures, and Columbia River flows, and consequently fish eggs and larvae distributions and survival. Long-term research is needed to identify how ichthyoplankton and fish recruitment are affected by regional and largescale oceanographic proces

Activity in the pallial nerve of knobbed (Busycon carica) and channeled (Busycotypus canaliculatum) whelks recorded during exposure of the osphradium to odorant solutions

Adult horseshoe crabs (Limulus polyphemus) are the preferred bait in the U.S. east coast whelk pot fishery, but their harvest is being restricted because of severe population declines in the Chesapeake and Delaware bays. To identify other baits, the activity in the pallial nerve of whelks was determined during exposure of the osphradium to odorant solutions prepared from horseshoe crab eggs, horseshoe crab hemolymph, and hard clam (Mercenaria mercenaria) tissue. All three elicited significant responses; bait based on them may provide an alternative to the use of adult horseshoe crabs, although extensive behavioral testing remains to be done. Channeled whelk did not respond to molecular weight fractions (>3 kDa and <3 kDa) prepared from horseshoe crab egg odorant solutions but did respond when the molecular weight fractions were recombined. Whelks appear to have broadly tuned chemoreceptors and manufactured baits may need to mimic the complex mixture of odorants derived from natural sources.

Timing and duration of mating and brooding periods of Atka mackerel (Pleurogrammus monopterygius) in the North Pacific Ocean

The timing and duration of the reproductive cycle of Atka mackerel (Pleurogrammus monopterygius) was validated by using observations from time-lapse video and data from archival tags, and the start, peak, and end of spawning and hatching were determined from an incubation model with aged egg samples and empirical incubation times ranging from 44 days at a water temperature of 9.85°C to 100 days at 3.89°C. From June to July, males ceased diel vertical movements, aggregated in nesting colonies, and established territories. Spawning began in late July, ended in mid-October, and peaked in early September. The male egg-brooding period that followed continued from late November to mid-January and duration was highly dependent on embryonic development as affected by ambient water temperature. Males exhibited brooding behavior for protracted periods at water depths from 23 to 117 m where average daily water temperatures ranged from 4.0° to 6.2°C. Knowledge about the timing of the reproductive cycle provides a framework for conserving Atka mackerel populations and investigating the physical and biological processes influencing recruitment.

Description of embryonic development of Atka mackerel (Pleurogrammus monopterygius)

Atka mackerel (Pleurogrammus monopterygius) is hexagrammid fish that inhabits the temperate and subarctic North Pacific Ocean and neighboring seas (Fig. 1). This highly abundant fish is a critically important prey species (Sinclair and Zeppelin, 2002; Zenger, 2004) that supports a directed commercial trawl fishery (Lowe et al., 2006). Atka mackerel is a demersal spawner and males provide parental care to eggs (Zolotov, 1993). During breeding periods, sexually mature males aggregate on the bottom at nesting sites where they establish territories (Lauth et al., in press). Sexually mature females periodically visit male nesting territories from July to October to spawn batches of demersal egg masses (McDermott and Lowe, 1997; McDermott et al., 2007). Individual nests may consist of multiple egg masses deposited by different females, and males defend nesting territories for a protracted period lasting from the time territories are being established until all eggs within the territory are completely hatched (Lauth et al., 2007). Knowledge about the timing of the reproductive cycle and the use of spawning habitat are important for understanding population structure and the dynamics of stock recruitment, which in turn are important factors in the management of Atka mackerel populations.

Diel variation in vertical distribution of an offshore ichthyoplankton community off the Oregon coast

We examined the diel ver-tical distribution, concentration, and community structure of ichthyoplank-ton from a single station 69 km off the central Oregon coast in the northeast Pacific Ocean. The 74 depth-stratified samples yielded 1571 fish larvae from 20 taxa, representing 11 families, and 128 fish eggs from 11 taxa within nine families. Dominant larval taxa were Sebastes spp. (rockfishes), Stenobra-chius leucopsarus (northern lampfish), Tarletonbeania crenularis (blue lan-ternfish), and Lyopsetta exilis (slender sole), and the dominant egg taxa were Sardinops sagax (Pacific sardine), Icichthys lockingtoni (medusafish), and Chauliodus macouni (Pacific viperfish). Larval concentrations generally increased from the surface to 50 m, then decreased with depth. Larval concentrations were higher at night than during the day, and there was evidence of larval diel vertical migration. Depth stratum was the most important factor explaining variability in larval and egg concentrations.